This is a revision of the completely limbate African Fissidens species with small. to medium sized, pluripapillose and mammillose laminal cells. These species belong to subg. Polypodiopsis Müll. Hal.) Broth. (formerly known as subg. Aloma (Kindb.) Pursell & Brugg.-Nann.) sect. Antennidens (Müll. Hal.) Paris (formerly known as sect. Semilimbidium Müll. Hal.). One new species, viz. F. latelimbatus Brugg.-Nann. sp. nov. is described here. Furthermore, a number of new synonyms is proposed. Thus, F. dupuisii Renauld & Cardot, F. dicranelloides Paris & Broth. in Paris, F. occultifolius Müll. Hal. ex Paris, F. circinicaulis Cardot, F. tisserantii Broth. & P. de la Varde in P. de la Varde, F. rugifolius Dixon, F. glauculus var. aculeifrons P. de la Varde in Thér. and F. undatus (Müll. Hal.) Par. are subsumed under F. dasyphus Welw. & Duby in Duby, while F. glaucissimus var. minor Welw. & Duby in Duby is subsumed under F. glaucissimus Welw. & Duby in Duby var. glaucissimus and F. glauculus var. gymnandrus P. de la Varde under F. unipapillosus Brugg.-Nann.. All species are provided with a newly confirmed geographic distribution. Country records not confirmed by me are provided with a reference. This paper further includes several new country records: Fissidens dasyphus is reported new to Sudan, Madagascar and Malawi, F. glaucissimus Welw. & Duby to Burundi, F. inaequalilimbatus Paris & Broth. to the Democratic Republic of the Congo, F. unipapillosus Brugg.-Nann. to Gabon and F. biformis Mitt. to Guinea. The following taxa are lectotypified: Fissidens dicranelloides Paris & Broth., F. dupuisii Renauld & Cardot, F. glaucissimus Welw. & Duby, F. glauculus var. aculeifrons P. de la Varde, F. g. var. gymnandrus P. de la Varde, F. inaequalilimbatus Paris & Broth., F. microphoenix P. de la Varde, F. occultifolius Müll. Hal. ex Paris (= Fissidens muelleri Dusén, hom. illeg.), F. rugifolius Dixon, F. subglaucissimus Broth., F. thallangae Dixon and F. tisserantii Broth. & P. de la Varde.
Introduction
This is the third part of a monograph of the Fissidentaceae of Tropical Africa. The first, Bruggeman-Nannenga (2013), deals with Fissidens Hedw. subg. Fissidens (i.e. completely limbate species with small to medium sized, smooth cells, bryoides-type peristomes and 40 or more exothecial cells on the capsule periphery). Though it focuses on the species from tropical eastern Africa, this paper covers most subg. Fissidens species from tropical western Africa as well.
The present paper is the second on subg. Polypodiopsis (Müll. Hal.) Broth., 1901. The first was published as ‘Fissidens subgenus Aloma (Bryophyta) in tropical Africa (Bruggeman-Nannenga 2017)’. Meanwhile it has appeared that the name Polypodiopsis has priority over Aloma (Kindb.) Pursell & Brugg.-Nann., 2004 on subgenus level. The 2017-paper revises the subg. Polypodiopsis species with very large cells. These species typically have smooth cells and vary from completely limbate to partly limbate and from costate to ecostate. The present paper deals with those subg. Polypodiopsis species that are limbate on all laminae, have small- to medium-sized, pluripapillose or mammillose cells and costate leaves.
The titles of these papers can be used as parts of an identification key. Each paper treats a group of species that share a set of easy to recognize characters. Thus, the species in Bruggeman-Nannenga (2013) have smooth cells and limbate, rarely semilimbate leaves, whereas the species in the 2017-paper have exceptionally large cells. However, a few species are ambiguous for the chosen characters. For instance, based on its medium sized, smooth, dorsal laminal cells the limbate F. zollingeri Mont., 1845 is treated in the 2013-paper, though its large vaginant laminal cells and sporophyte suggest that it is closer related to the species in the 2017-paper. Similarly, in the present paper, F. dasyphus and F. latelimbatus can be limbate or semilimbate. These species will, therefore, be treated again in the paper on semilimbate species.
Worldwide only a few species have limbate leaves in combination with small- or medium-sized pluripapillose or mammillose cells. Apart from the species treated here, there are only the neotropical F. goyazensis Broth., 1895, F. yucatanensis Steere, 1935, F. weirii Mitt., 1869 and F. plurisetus E.B.Bartram in B.Willis, 1939, the Asian F. benitotanii Z. Iwats., K.T.Yong & Tad. Suzuki, 2009 and the Polynesian F. raiatensis E.B.Bartram, 1931.
Material and methods
The arrangement in the distributions follows Hollis and Brummitt (1992). All distributions are updated and when necessary emended. Distribution data for Africa without reference have been confirmed by specimens seen for this paper.
In my earlier papers measurements were made with an eyepiece with graticule. In more recent ones measurements of leaves, spores and cells are made from photographs with the program piximètre.
Papillae are observed in water and/or in 5% KOH and photographed.
Measurements
Stem length is measured without proliferations.
Laminal cells. The unique morphology of the leaf has important consequences. Most importantly, cells of vaginant lamina cells differ essentially from those of the apical and dorsal lamina as they have papillae on the exterior side only. Consequently, vaginant laminal cells are, particularly in pluripapillose species, easier to analyze. When making measurements one should further be aware that within a leaf there can be considerable differences in shape, size, papillae and diameter of the walls. Cells close to stem-, costa – and limbidium tissue tend to be larger and more elongated and often have thicker, less papillose walls than the inner laminal cells. To obtain a high level of comparability between measurements in different publications, researchers should make sure to measure the same cells. In my later papers measurements are from mid vaginant laminal (Fig. 1A) and mid dorsal laminal cells (Fig. 1B). In my earlier papers cell length and width included the walls, in my more recent papers the walls are excluded.
Papillae. Position, shape and number of papillae are important for distinguishing species. Unfortunately, papillae can be quite variable, even in one and the same stem. This is probably due to environmental factors and age. This variability makes papillae a challenging diagnostic trait. Part of this problem can be eliminated by standardized observations. Papillae are best analyzed in the mid vaginant lamina (Fig. 1A) in 5% KOH. As they often appear different at different focusing levels a standardized way of measuring is needed. My measurements are from the surface level of papillae on the mid laminal cells.
Terminology
Papillae can be ‘over the lumen’ (Fig. 2H), marginal (also indicated as ‘at the corners’) (Fig. 3I), or anomalous. Typically, papillae are easily identified as either ‘over the lumen’ or marginal, but this is not always the case.
Leaf and laminae. The Fissidens leaf is unique in having a duplicated part that embraces the stem, the so-called vaginant lamina and two simple laminae growing from the costa. The abaxial dorsal lamina extending the whole length of the leaf and the apical lamina distal of the vaginant lamina (Fig. 1).
Mammillose. No distinction is made between mammillose and unipapillose.
Completely limbate is used when limbidia are present on the vaginant, apical and dorsal lamina regardless of whether the limbidia reach the apex and the insertions.
Key
Pluripapillose species
Fissidens dasyphus Welw. & Duby (Fig. 2)
In Duby, Mém. Soc. Phys. Genève 21: 224, tab. 4, f. 2, 1872 – Protologue: [Angola], in argillaceis praeruptis montium circa Sange et Bumba a 2000–2200 ped. altit. praesidii Golungo, Welwitsch – lectotype (designated by Bruggeman-Nannenga and Pursell 1995): Welwitsch Iter Angolense 137 (BM).
Conomitrium undatum Müll. Hal., Linnaea 39: 362. 1875; Fissidens undatus (Müll. Hal.) Par., Index Bryol. 489, 1869 – protologue (two syntypes): [Sudan, see remark on Schweinfurth's collections below], Bongo-regiones, in parietibus inundatis saxosis ferruginosis (Raseneisenstein) cryptae ad Gubbihih prope Kulongo, 31 Julio 1896, Schweinfurth s.n.; Djur-regiones [Jur], Seriba magna Ghattas 1869, Schweinfurth s.n.(not seen) – lectotype (designated by Bruggeman-Nannenga 1997) labelled: Central Afrika, Lande der Bongo [SUDAN], in parietibus inundatis speluncae prope Gubbihih circa Kulongo, 31 VII 1869. Schweinfurth s.n. (BM-herb. Bescherelle; isolectotypes: BM (herb. Hampe. 2 collections); PC-PV as Schweinfurth 2333C (MNHN-PC-0096289) and H-BR as Schweinfurth 2333). syn. nov.
Fissidens glauculus Müll. Hal. ex Dusén, Kongl. Svenska Vetensk. Acad. Handl., n. s. 28: 12a, tab. 2, f. 2, 1895 – protologue (three syntypes): 1) Camerunia, in formicetis, prope Isongo pagum, II 1892 c. fr., Dusén; 2), Camerunia, ad Ekundu Ndene, III 1892, Dusén; 3) Camerunia, ad Ndian emporium, III 1892 c. fr., Dusén – lectotype (designated by Bruggeman-Nannenga 1997): Musci Africani in Camerunia a P. Dusén collecti 760. Ad N' dian emporium, III 1892 (H-BR; isolectotype: PC).
Fissidens dupuisii Renauld & Cardot, Bull. Soc. R. Bot. Belgique 35: 306, 1897; F. glauculus subsp. dupuisii (Renauld & Cardot) P. de la Varde, Ann. Cryptog. Exot. 2: 286. 1929; F. glauculus var. dupuisii (Renauld & Cardot) P. de la Varde, Mém. Soc. Sci. Nat. Cherbourg 42: 28, 1936 – protologue: [Democratic Republic of the Congo], Africa tropicalis, in silva Bidi, ditione fluminis Congo, Dupuis – lectotype (designated here): Afr. trop., Congo belge, forêt de Bidi, Dupuis 486 (MNHN-PC-PC0130827); isolectotypes: PC, H-BR. syn. nov.
Fissidens dicranelloides Paris & Broth. in Paris, Rev. Bryol. 31: 42,1904; F. glauculus subsp. dicranelloides (Paris & Broth.) P. de la Varde, Ann. Cryptog. Exot. 2: 286. 1929 – protologue: [Guinea], ad arbores prope pagum Dieudieu, cercle de Kouroussa, VII 1903. Pobeguin s.n. – lectotype (designated here): [Guinea] Guinée française. cercle de Kouroussa, prope pagum Dieudieu. Julio 1903, Pobeguin s.n. (MNHNPC-PC0106722; isolectotypes: H-BR, PC). syn. nov.
Fissidens occultifolius Müll. Hal. ex Paris, Index Bryol. (ed. 2) 2: 217, 1904, nom. nov. for Fissidens muelleri Dusén, Kongl. Svenska Vetensk. Acad. Handl., n. s. 28 (2): 14, a, tab. 2 f. 9. 1895 hom. illeg.; Conomitrium occultifolium Müll. Hall., Gen. Musc. Frond. 74. 1900 nom. nud. – protologue: Africa, in territorio Oil Rivers in humo, ad Old Calabar emporium, VII 1892, Dusén – lectotype (designated here): [NIGERIA], West Africa, Old Calabar, on the ground, 28 VII 1892, Dusén s.n. (H-BR; isolectotypes: PC-PV as C. occultifolium, Musc. Afr. a P. Dusén coll. 905 and 906) syn. nov.
Fissidens circinicaulis Cardot, Rev. Bryol. 35: 65, 1908; F. glauculus var. circinicaulis (Cardot) P. de la Varde, Ann. Cryptog. Exot. 2: 284, 1929 – protologue: [Senegal], Casamance, Oussouye, sur la terre, M. et Mme Ch. Matthieu – lectotype (designated by Bruggeman-Nannenga 1997): [Senegal], Casamance, Oussouye, à terre, 1907, M. et Mme Ch. Mathieu), sp. nova (MNHN-PC-PC0095416; isolectotypes: PC-PV and H-BR). syn. nov.
Fissidens tisserantii Broth. & P. de la Varde in Potier de la Varde, Bull. Soc. Bot. France 72: 793. 19 f. 5, 1925 – protologue: [Central African Republic], 3 syntypes, leg. R.P. Tisserant: 1) Sur termitières près de Boedou, 2) près d'Ippy, 3) terrains argileux, près Moronbas (not seen) – lectotype (designated here): Central African Republic, [Oubangui], Riv. près Baedou, sur termitière; sous-bois, 29 V1294. R.P Tisserant s.n. (H-BR; isolectotype PC-PV as Tisserant 7). syn. nov.
Fissidens rugifolius Dixon, J. Bot. 76: 220, Fig. 18, 1938 – protologue: country not indicated, primary forest, 1200 m, on ant hill, 30 VI 1935, A.S. Thomas 1359 – lectotype (or holotype?) designated here): [Uganda, distr., Sesse Co.], Towa Forest, Bugala [Island], on anthill, short dark moss, 4000 ft; 30 VI 1935, Thomas 1359 (BM 000 726). syn. nov.
Fissidens glauculus var. aculeifrons P. de la Varde in Thér., Bull. Mus. Natl. Hist. Nat., Sér. 2, 3: 771, 1931 – protologue: Guinea [Guinée française], Conakry, en compagnie de l'espèce précédente [F. muelleri], Chevalier s.n., 1930 – lectotype (designated here) Guinee Française, Conakry, XI 1930. Chevalier s.n. (MNHN-PC-PC0106727; isolectotypes: PC). syn. nov.
Misapplied name: Fissidens weirii sensu Bruggeman-Nannenga and Pursell (1995), Bruggeman-Nannenga (2009). It appeared that African specimens are F. dasyphus and neotropical ones F. weirii Mitt. s.s. (Diop et al. 2018).
Rectification: the specimen cited by Bruggeman-Nannenga (1993) as type of F. glauculus, viz. Guinea, Conakry, Chevalier s.n., is not the type of F. glauculus, but that of F. glauculus var. aculeifrons. This error has no consequences for the synonymy since F. glauculus C.Müll. var. aculeifrons is no more than a narrow-leaved, freakish expression of F. dasyphus.
Remarks
Fleischer (1904) and O'Shea (2006), incorrectly considered the pluripapillose F. occultifolius Müll. Hal. ex Paris (= F. muelleri Dusén hom. illeg.) to be a synonym of F. schmidii Müll. Hal., 1853 (= F. crispus Mont. 1838). Fissidens crispus has smooth cells and belongs to subg. Fissidens. Bruggeman-Nannenga (1993) synonymized F. glauculus and F. muelleri Dusén (this should have been with F. occultifolius since the illegitimate homonym F. muelleri Dusén had been replaced by that name).
The synonymy of F. circinicaulis, F. dupuisii, F. dicranelloides, F. occultifolius (F. muelleri Dusén hom. illeg.), F. rugifolius, F. tisserantii follows from earlier reductions to F. weirii and F. glauculus respectively (Potier de la Varde 1929, Bruggeman-Nannenga 1993, 1997, 2009, Bruggeman-Nannenga and Pursell 1995). Fissidens undatus is a semilimbate expression that was earlier considered to be F. intramarginatus (Hampe) A. Jaeger, 1869 (Bruggeman-Nannenga 1997).
Diagnosis and variability
Fissidens dasyphus is characterized by oblong, often undulate leaves, dorsal lamina and vaginant laminae rounded at the insertion; vaginant laminae are about as wide to slightly wider than the stem, limbidia narrow and marginal throughout, reaching the insertion of the vaginant lamina and laminal cells with 1–3 (–5) papillae over the lumen. Archegonia typically terminal on main shoots and branches, but solitary axillary archegonia occasionally occur (e.g. in type of F. tisserantii), perigonia terminal on branches, in one case (Heras 522/94) clusters of naked axillary antheridia in upper leaves of perichaetial stems were observed.
Fissidens dasyphus varies from limbate on all laminae to semilimbate (limbate on the vaginant lamina only) (Potier de la Varde 1929, 1936, Bruggeman-Nannenga 1993). Purely limbate, purely semilimbate expressions and mixed collections are all common. In limbate specimens, ‘e.g. the type specimen of F. dasyphus’, limbidia end below the leaf apex and above the insertion of the dorsal lamina. Limbate expressions have been described as e.g. F. dasyphus and F. glauculus, transitional ones, i.e. semilimbate specimens with poorly developed limbidia on some dorsal lamina, as F. dupuisii and F. dicranelloides and semilimbate ones as F. undatus and F. tisserantii. It appeared to impossible to find a combination of at least two traits by which these expressions can be distinguished. Thus, they are considered to constitute one variable species, F. dasyphus. Limbate expressions are most likely to be confused with F. glaucissimus and F. inaequalilimbatus. For differences see key. Semilimbate expressions are recognized by the combination of oblong, often undulate leaves (undulations most easily seen in 5% KOH), narrow, marginal limbidia extending from the insertion to apex of the vaginant laminae and laminal cells with 1–3 (–5) papillae over the lumen. They can be confused with F. intramarginatus s.l. This species complex requires revision.
Description
Stem in cross-section with central strand, 3.0–7.0 × 1.2–2.0 mm, unbranched or branched from old perichaetia, hardly heterocaulous, with close to distant, pinnately arranged leaves; rhizoids yellowish brown, less often dark brown, smooth; axillary nodules and cells not differentiated; leaves pale green, 15–17 pairs, crispate when dry, often undulate, oblong to oblong-lanceolate leaves with wide acute to acute-acuminate apex, (0.9–) 1.2–1.9 × 0.2–0.3 (–0.4) mm, (3.6–) 4.0–6.5 times as long as wide, limbate; limbidium marginal, typically present on all laminae of mid and upper leaves, in lower leaves frequently poorly developed, ending distinctly above the insertion of the dorsal lamina and below the leaf apex, reaching insertion of vaginant laminae, typically confluent at the apex of the vaginant lamina (often hard to observe); in mid dorsal lamina 5–8 µm wide and bistratose, on vaginant laminae up to five cells wide, 11–24 µm wide and uni- to 2 (–4) -stratose; vaginant lamina 3/5–3/4 the leaf length, at the insertion rounded, about as wide as to slightly wider than the stem, margins often reflexed, subequal; dorsal lamina rounded below, reaching the insertion, not decurrent; dorsal and apical lamina unistratose; costa per- to excurrent, occasionally ending two cells below the apex, bryoides-type; mid dorsal laminal cells 4.5–8.0 × 2.5–7.5 µm, with 1–3 (–4) papillae over the lumen, often seemingly smooth; mid vaginant laminal cells oblong, firm-walled, 4.0–8.5 × 3.0–6.0 µm with 1–3 (–5) papillae over the lumen (papillae often hard to observe). Axillary gemmae (Fig. 2L, Heras 508/94) and tubers (Richards 3513) rare. Perigonia and perichaetia on the same stem, perigonia terminal on 1.0–4.5 mm long stems, less often on budlike axillary branches (1–2 per axil), or in naked, axillary clusters, perigonial branches frequently growing from old perichaetia; antheridia 160–210 µm long; perichaetia terminal on plants and branches, perichaetial branches often growing from old perichaetia, archegonia 280–300 µm long; perichaetial leaves 1–2 mm long; calyptra scabrous, 0.5 mm long. Sporophyte, seta 3–4 mm long, smooth, 1–2 per perichaetium; capsule cylindrical to gibbose, 0.75 × 0.35 mm wide, with 32 columns of ± isodiametric, collenchymatous exothecial cells; peristome scariosus type, tooth base 32 µm wide; operculum long rostrate, 0.55–0.6 mm long; spores subglobose, (6.0–) 9.0–10.5 (–11.5) µm diam, finely papillose, green.
Substrate and habit
Most collections are from forests (lowland-, submontane-, montane- and summit-, rain-, riverine-, degraded). Occasionally reported from other habitats e.g. farmland and a rubber plantation. Most often on termite mounds and soil, less often lignicolous (on bark, epiphytic, on base of Raphia palm and decaying wood) or rupicolous. Growing in mats. Altitude: 75–2300 m.
Distribution
A common, wide-spread African endemic, known from West Tropical Africa (Benin, Burkina Faso (O'Shea 2006), Ghana, Guinea, Ivory Coast, Liberia, Nigeria, Senegal, Sierra Leone, Togo (O'Shea 2006); West-central Tropical Africa (Cameroon, Central African Republic, Congo, Democratic Republic of the Congo, Equatorial Guinee (Bioko, Rio Muni), Gabon); North East Tropical Africa (Sudan, see remark below); East Tropical Africa (Kenya, Tanzania, Uganda); South Tropical Africa (Angola, Malawi) and Western Indian Ocean (Madagascar) – New to Madagascar and Malawi. The Malawi-specimen was earlier cited as F. intramarginatus (O'Shea et al. 2001). New to Sudan under this name (type of Conomitrium undatum Müll. Hal.).
Note
When Schweinfurth made his expedition to Central in Africa (1868–1871) the European colonization had hardly started. Since the present geography of Africa for a large part reflects the borders of former colonies and is not easy to correlate Schweinfurth's labels with present countries. For instance, the syntypes of F. undatus have been collected in 1869 in the Bongo- and Djur-regiones respectively. A reprint of a map of Africa printed in 1877 (Anstadt et al. 1992) locates both the Bongos and the Djur (as Djoer) in what is now Sudan. This is supported by Wickens (1972) who describes Schweinfurt's travels in great detail and shows that Schweinfurth visited southern Sudan from 1 IV 1869 to 17 IX 1869.
Selected specimens
Angola, 21 miles West of Vila Salazar, Brass 21000 (EGR); – Benin, Atacora, Kota (Natitingou), 10°12′63.3″N, 01°26′.6″E. B. Alia AM636 (HNB, private herbarium Bruggeman-Nannenga); – Cameroon, Lolodorf, Standt 251 (H-BR); – Central African Republic, près d' Ippy, Tisserant 19 (H-BR, PC); Galerie près Zubingé [Zubuyé], 25 km S.O. Ippy, Tisserant 104 (PC-PV, H-BR, c. spor.); – Congo, bas fond de Getamayanon, Assel 743 (PC-PV); – Democratic Republic of the Congo, Prov. Kivu, Pinga, 96 km NW Goma. Flusstal 4 km NNO Ort. östl. Ort. 1°0′S, 28°43′E, Müller Z 226 (DR, L); Pinga, 96 km NW Goma. Flusstal 4 km NNO Ort. östl. Ort. 1°0′S, 28°43′E, Müller Z 232 (DR, L); Prov. Equateur, Kalamba, 55 km S Mbandaka 3 km östl. Ort. 0°25′S, 18°19′E, Müller Z 235 (DR, L); Irangi Forest Station 110 km W of Bukavu. Summit Forest of Mt Ilimo, Bryotrop Exp. 1991: Frahm 6857 (private herb. Bruggeman-Nannenga) and Pócs 6780 (private herb. Bruggeman-Nannenga); Haute-Zaïre, route Kisangani-Ubundu (au S de Kisangani), Lisowski 50557 (Kram, L); – Equatorial Guinea, Insel Bioko (Fernando Po), Berg Caldera 10 km S Luba, am NON-Hang am Weg von Ruiche zum Gipfel. 3°24′N, 8°33′E, F. Müller B 101 (DR, L); Muni, Evinayong, Monte Alen. Moka, Lejoly's transect. Paxti Heras 387/94 (ALAVA, L); Niefang, Monte Chocolate, Heras 508/94 (Alava, L); Niefang, Monte Chocolate, Heras 522/94 (Alava, L); – Gabon, Prov. Ogooue-Ivindo, Makokou, Réserve Intégrale d'Ipassa, Makokou, station de recherche d'Ipassa, 0°30′42′N, 12°48′12′E, Vanderpoorten GAB1237 (L); – Ghana, Eeastern Region, Brukru Rock, Kwahu Tafo, Jones 1418 (BM, L); Ashanti Region, Bobiri Forest Reserve. 6.41°N, 1.20°W, J. Rikkinen 040001-24 (H, L); 26 km NW from Kumasi, Kovács 18a (PC); – Guinea, nr Macenta, in ‘Magnetic Mountain’, Lisowski 196 (Kram) and 213 (Kram, L); – Ivory Coast, Umgebung der Ortschaft Gouedie (= Gouele), 24 km NW Man, 7°32′N, 7°45′W, Müller E 206A (DR, L); – Kenya, north Kavirondo District, S. E. of road between Rondo and Kakamega Sawmills, earth-covered rock in flash stream bed in primary forest, Townsend 77/325A (E); – Liberia, Gbanga. IX 1926 Linder s.n. (PC-PV); Madagaskar, Prov. Antsiranana, Parc National de la Montagne d’Ambre. 12°30′S, 49°10′E. Open mountain forest. Touw & Snoek 25099 (L); – Malawi, Mulanje, Mulanje Mt Muluzi River, NGHodgetts 2011 (E); – Nigeria, Ijebu-Ode Prov., Shasha Forest Reserve. Near Akilla. Richards 3513 (PC-PV); Zaria Prov., Anara Forest Reserve, Keay 13491 (MO, L); Colony Province, Epe, McFarlane 13 (L); Western Region, Ijebu Prov., Ijebu-Ode, Idowa, McFarlane 521 (L); – Senegal, type specimen F. circinicaulis; Sierra Leone, Kangari Hills, 1 km NW of Masuri, Marmo s.n. (H, L); – Sudan, lectotype F. undatus; – Tanzania, West Usambara Mts, near Mazumbai village, along road leading northwards from the Field Station, Pócs 8540/S (EGR, L); Nguru Mts in Morogoro District. W of Magole Village, Pócs & Minja 88182/A (EGR, L); – Uganda, Kalangala District, Ssese Islands, Bugala Island, at Mkalwe spring near Kalangala village, 00°19′3″S, 32°15′E, Pócs, Lyle & Samuela 97105/E (EGR, L).
Illustrations
Bruggeman-Nannenga (2006b): Fig. 24f–k as F. weirii; Chuah-Petiot 2003: Fig. 36 as F. weirii; Dixon 1938: Fig. 18a as F. rugifolius; Dusén 1895: 12, a, tab. II, f. 2 as F. glauculus and 14a, tab. II f. 9 as F. muelleri; Ngaba Zogo 1983: Fig. 2B as F. glauculus; Potier de la Varde 1931: Fig. 11: 7 as F. tisserantii and Fig. 12: 7 as F. undatus.
Figure 2.
Fissidens dasyphus. (A) vegetative stem, (B) sporophytic stem, (C–D) leaves, (E) leaf apex, (F) mid leaf, (G) basal part of leaf, (H) cells of vaginant lamina, (I) limbidium of mid dorsal lamina (poorly limbate specimen), (J) peristome tooth, (K) capsule with operculum, (L) gemmae. (A, G) from Lisowski 196, (B) from Tisserant 104, (C, E–F) from F. Müller Z232, (D) from Pócs 6780, (H) from F. Müller Z235, (I) from Vanderpoorten GAB 1237, (J) from Kovács 18A, (K) from Frahm s.n. and (L) from Heras 508/94.
Fissidens glaucissimus Welw. & Duby (Fig. 3)
In Duby, Mém. Soc. Phys. Genève 21: 223, T. 3, f. 4a–e, 1872 – protologue: syntype 1: [Angola], in cryptis maxime umbrosis rupium montium editiorum (2400 ped. alt.) de Quilongo in praesidio Golungo, Welwitsch; syntype 2: in rupium excavationibus aestate inundatis juxta ripas fluminis Luinho intra faciem internam cryptarum denso agmine viroris glauci decoratam, Welwitsch – lectotype (designated here) labeled: Welwitsch, Iter Angolense 142. Angola, Golungo Alto. Mont. de Queta ad Flumin. Luinha. In cryptis rupium schistos. sylvar. primit. (BM; isolectotypes with slightly different labels: BM, PC and H-SOL).
Fissidens glaucissimus var. minor Welw. & Duby in Duby, Mém. Soc. Phys. Genève 21: 224, 1872 – protologue: Angola, ad rupes gypsaceas muscosas juxta rivulos aut in sylvis – considered a synonym because the original description fits within the variability observed in this species. syn. nov.
FissidenssubglaucissimusBroth.,Bol.Soc.Brot.8:175,1890 – protologue: INS. S. THOMÉ, ad corticem arborum, Fr. Quintas – lectotype (designated here): Ins. S. Thomé, 170 m, Quintas 5 (H-BR; isolectotype: MNHN-PC-PC0106769).
Fissidens microphoenix P. de la Varde, Bull. Soc. Bot. France 74: 142, f. 39, 1927; F. subglaucissimus Broth. var. microphoenix (P. de la Varde) P. de la Varde, Mém. Soc. Sci. Nat. Math. Cherbourg 42: 27, f. 6, 1936 – lectotype (designated here): GABON, chute de la Macenga près Dibwangui, sur rochers feldspatiques, Le Testu 5626 (MNHN-PC-PC0106266, isolectotype: MNHN-PC-PC0106267).
Fissidens thallangae Dixon, J. Bot 76: 220, 1938; F. skallangae Dixon ex Wijk, Margad. & Florsch., Regnum Veg. 26: 315. 1962. (orthograpic error for F. thallangae) – lectotype (or holotype?) (designated here): Uganda, Thallanga Forest, Imatong Mts, R. Loboli: damp ground, common moss, 2950 ft, 16 XII 1935, A. S. Thomas 1576 (BM).
Diagnosis
Fissidens glaucissimus is readily recognized by its stiff, obscure, linear, limbate leaves, narrowly acute apices, vaginant laminae reaching ± 3/4 the leaf length, the exceptionally wide, unistratose limbidia of the vaginant lamina and vaginant laminal cells 4.0–7.5 (–8.5) µm long with 7–10 (–12) marginal papillae. The rhizoids are frequently red, but like in other species with red rhizoids, this is not a constant character. The species is most likely to be confused with F. inaequalilimbatus, which differs by narrower limbidia (up to six rows of cells) on the vaginant laminae and mostly wider leaves. Linear, obscure leaves are also characteristic of the Asiatic species F. wichurae Broth. & M. Fleisch., 1899, F. speluncae Broth. in Herzog, 1910 and F. kinabaluensis Z. Iwats., 1996, but these species are semilimbate.
Description
Stem in cross-section with central strand, unbranched or branched, branches often sprouting from old perichaetia, pinnately foliated, hardly heterocaulous; rhizoids red, red-brown or brown, smooth; axillary nodules and cells not to slightly differentiated; stem 6.0–12.0 × 1.5–3.0 mm, leaves dark green, close, 16–34 pairs, hardly altered when dry, linear to linear-lanceolate, narrowly acute to acute-acuminate, 1.00–2.10 × 0.15–0.30 mm, (5–) 6–8 times as long as wide, margin limbate, crenulate where elimbate; limbidia on all laminae of all leaves, occasionally branched, reaching to almost reaching the leaf apex, fusing at the apex of the vaginant lamina, reaching the insertion of the vaginant lamina, ± reaching the insertion of the dorsal lamina to ending well above it, in mid dorsal lamina 5–9 µm wide, marginal, in cross-section thinner than inner laminal cells, unistratose; limbidium of proximal vaginant laminae (6–)7–14 cells and 35–65 µm wide, marginal, unistratose, in cross-section equally thick as inner laminal cells; vaginant lamina 7/10–3/4 the leaf length, in basal part narrower than to about as wide as the stem (in perichaetial stems sometimes slightly wider than the stem), straight towards the insertion or strongly widened and then concave towards the insertion, unistratose, unequal with the smaller lamella ending closer to the margin than to the costa; dorsal lamina narrow, straight to slightly rounded towards the insertion, occasionally basal most part reduced to limbidial tissue, mostly reaching the insertion; mid dorsal laminal cells obscure, (3.0–) 4.0–7.5 × 3.0–5.5 µm with 6–10 marginal papillae; mid vaginant laminal cells 4.0–7.5 (–8.5) × (2.0–) 3.0–5.5 (–6.5) µm with 7–10 (–12) marginal papillae. No gemmae seen. Perichaetia and perigonia on the same or separate stems; antheridia terminal on tiny, 0.6–1.8 mm long stems at the base of perichaetial stems and on ± 0.5 mm long, bud-shaped perigonial branches; antheridia 150–250 µm long; perichaetia terminal on plants or, less often, branches, perichaetial branches from upper and middle part of stem; perichaetial leaves 1.8–2.3 mm long, archegonia (250–) 350–470 µm long; calyptra 0.75 mm long, smooth. Sporophyte, seta 3.0–7.5 mm long, smooth, 1 (–2) per perichaetium; capsule gibbose, 0.8–1.2 × 0.4–0.6 mm, ± 32 (–42) columns of quadratic, collenchymatous exothecial cells; peristome scariosus-type, tooth base 43.5–50.0 µm wide; operculum not seen; spores subglobose to ellipsoid, 7.5–12.0 µm long, smooth to finely papillose.
Substrate and habit
Mostly on rock and soil, occasionally on termite nests, rarely on wood. Growing in mats, in forests, on banks near sources and on in rivulets. Altitude: sea-level–1500 m.
Distribution
Endemic to tropical Africa. Known from: West Tropical Africa (Guinea, Ivory Coast (O'Shea 2006), Ghana (Hodgetts et al. 2016), Nigeria, Togo); West-Central Tropical Africa (Cameroon, Central African Republic, Democratic Republic of the Congo, Equatorial Guinea (Bioko & Muni), Gabon, Sao Tomé & Principe); East Tropical Africa (Burundi, Kenya (Chuah-Petiot 1996), Tanzania and Uganda) and South Tropical Africa (Angola) – Common in West Central, South Tropical Africa and East Tropical Africa, but apparently rare in Tanzania where in spite of extensive collecting it has been collected only twice – New to Burundi.
Selected specimens
Angola, lectotype F. glaucissimus; – Burundi, Territoire de Bururi, près de la Mission de Mugara, De Sloover 19.241(PC, L); – Cameroon, Solodorf, Standt 112 (H-BR); Südkamerun, Waldgebiet, Bezirk Ebolowa, Mildbraed 5759; – Central African Republic, Plateau de Kaga-Dasuba, Assel 722 (PC); Bangui, colline derrière le camp de Kasaï, Assel 116 (PC); – Democratic Republic of the Congo, Prov. Kivu, 96 km NW Goma, Flusstal 4 km NNO Ort. östl. Ort. 1°0′S, 28°43′E, Müller Z 219 (DR, private herb. Bruggeman-Nannenga); Irangi Forest Station 110 km W of Bukavu, Bryotrop Exp. 1991: Pócs 6757 (private herb. Bruggeman-Nannenga); UELE, Zone de'Aketi, 5 km à l'W de Likati. Lisowski 46454 (Kram, L); Haute-Zaïre, Ituri, env. de Nduye, Lisowski 43583 (Kram, L); – Equatorial Guinea, Bioko, Ureca W, Küste zwischen der Mündung des Rio Ole und Punta Sagre (Playa de Moraca); 3°16′N, 8°29′E, Müller B 426 (DR, L); Muni, Evinayong, Monte Alen. Moka, Lejoly's transect, Paxti Heras 381/94 (Alava, L); – Gabon, Ogooué-Ivindo, Makokou, 0°30′43.0′N, 12°48′12.1′E, Vandenpoorten GAB1699 (L); – Guinea, Region de Nzerekore, près de Macenta, Lisowski 964 (Kram, L); – Nigeria, Western Region, Ijebu Province, Ijebu-Ode, Odosenlu Alaro, McFarlane 558 (private herbarium Bruggeman-Nannenga); Prov. Calabar, Oban, cutline W of mile 51, Richards 5155 (PC-PV); Benin Prov., Div. Benin, Okomu Forest Reserve, Richards 3731 (PC-PV); Sao Tomé & Principe, Ins. Principis, Quintas 3. (H-BR: paratype F. subglaucissimus); Insula Sao Tome, Quintas 5 (H-BR lectotype F. subglaucissimus); – Tanzanina, Ost Usambari, Amani, Brunnthaler s.n. (H-BR); – Togo, Stadiaburg, 23 XII 1890, Büttner (H-BR); – Uganda: Budongo Forest, Nyakafunjo Nature Reserve, 1°42′ 50′N, 31°31′16′E, M.J. Wigginton U 3119A (E, L); Hoima District, Bugoma Forest Reserve, 12 km SE from Lake Albert, 4-6 km W of Kabwoya village; 1°14.2′N, 31°2-3′E; Lye & Pócs 97122/O (L, EGR).
Illustrations
Bruggeman-Nannenga 2006a: Fig. 10; Potier de la Varde 1936: Fig. 6 as F. subglaucissimus var. microphoenix; Ngaba Zogo 1983: Fig. 2A.
Figure 3.
Fissidens glaucissimus. (A) sporophytic stem, (B) perichaetial stem, (C) perigonial branch, (D–E) leaves, (F) leaf apex, (G) mid leaf, (H) insertion leaf, (I) papillae mid vaginant laminal cells, (J) cross-section leaf (high in vaginant lamina, dorsal lamina broken off), (K) cross-section limbidium (low in vaginant lamina), (L) cross-section stem. (A) from Müller Z 219, (B–C) from Lyle & Pócs 97122/O, (D, K, L) from VandePoorten GAB 1699, (E, I) from Müller B426, (F) from Mildbraed 5759, (G–H) from Pócs 6757, (J) from Heras 381/94.
Fissidens inaequalilimbatus Paris & Broth. (Fig. 4)
In Paris, Rev. Bryol. 31: 118, 1904 – protologue: Guinée française, ad ripas amnis Dieudieu. Julio 1903. Pobeguin s.n. – Lectotype (designated here): [Guinea], Guinea gall., cercle de Kouroussa, ad amnem Dieudieu, H . Pobeguin 83 (H-BR; isolectotype in PC-PV).
Diagnosis
Fissidens inaequalilimbatus is characterized by rather short, 0.8–1.0 mm long, lanceolate leaves that are limbate on all laminae of all leaves, pluripapillose cells and marginal papillae. It resembles F. glaucissimus from which it can be distinguished by the narrower limbidia and leaves that are typically wider.
Description
vegetative parts based on type-specimen, fertile parts on Hepper 7620).
Yellowish-orange in KOH, stem unbranched or with proliferations from old perichaetia, pinnately foliated, hardly heterocaulous, without central strand, 4.0–7.0 × 1.0–1.5 mm; rhizoids brown, smooth; axillary cells slightly differentiated, not protruding; leaves close to distant, up to 24 pairs, somewhat crisped when dry, lanceolate with acute-mucronate to rounded obtuse-mucronate apex, 0.8–1.0 × 0.2–0.3 mm, 3.1–3.8 times as long as wide, margin limbate, crenulate where elimbate; limbidia on all laminae of all leaves nearly or completely reaching the leaf apex, fusing at the apex of the vaginant lamina, reaching the insertion of the vaginant lamina, ending well above insertion of dorsal lamina, on the mid dorsal lamina 5–6 µm wide, marginal, unistratose; on the vaginant laminae 5–6 cells (sometimes more on perichaetial leaves) and 16 µm wide, unistratose, marginal or intramarginal; vaginant lamina 3/4 the leaf length, at the base about as wide as the stem, rounded towards the insertion, unistratose, subequal; dorsal lamina slightly rounded below, reaching the insertion, not decurrent, dorsal and apical lamina unistratose; costa excurrent, bryoides-type; mid dorsal laminal cells obscure, 4.0–7.0 × 3.0–4.6 µm, with 6–7 (–10) marginal papillae; mid vaginant laminal cells 6.0–12.5 (–14.0) × 4.0–9.5 µm with 6–12 as a rule marginal papillae. Gemmae not observed. Perigonia sparse, terminal on short, ± 1 mm long, branches; antheridia 170–220 µm; perichaetia terminal, rarely with a few caulogenous, solitary archegonia; perichaetial leaves 1.25 mm long, archegonia 210–250 µm long. Sporophyte not observed.
Substrate and habit
On various substrates. On soil, rocks, termite mounds and base of Elaeis-stem. Growing in mats or scattered between other mosses. In forest, often on banks, frequently near rivers. Altitude: 420–1260 m.
Distribution
Endemic to West Tropical Africa (Ghana, Guinea, Nigeria) and West Central Tropical Africa (Central African Republic, Democratic Republic of the Congo). Uncommon.
Selected specimens
Central African Republic, Bangui, Colline de Oubangui, 4°16′N, 8°49′E, Prendergast 145 (L); près Bangui, rochers de la fleuve, 500 m, Balázs 88/C (PC, EGR, L); Damara, forêt du Lac Sorcier, Assel 161 (PC); – Democratic Republic of the Congo, Kisangani, Plateau médical, base d' un tronc d' Elaeis guineensis. Lisowski 46405 (KRAM, L); – Ghana, Eastern Region, Akosombo, Hepper 7620 (L, E); Southern Scarp FR, road between Nkakaw and Mpraeso; 6°33′N, 0°40′W; Richards 7085 (MO mixed with F. schweinfurthii and other Fissidens species); – Guinea, District Kankan, bank of River Milo, near village Bordo, Lisowski 7236 (Kram, L); Conakry, Chevalier 1A (PC, mixed with type of F. latelimbatus); – Nigeria, Vom, Benue Plateau State, 8 May 1966, McFarlane 672, mixed with other Fissidens species (private herbarium Bruggeman-Nannenga).
Figure 4.
Fissidens inaequalilimbatus. (A–B) vegetative stems, (C–D) leaves, (E–F) leaf apices, (G) mid leaf, (H) insertion leaf, (I) cells mid vaginant lamina, (J) cross-section leaf. (A) from Prendergast 145, (B) from Assel 161, (C, E, H, J) from Hepper 7620, (D, F, G, I) from Lisowski 46405.
Fissidens latelimbatus sp. nov. (Fig. 5)
Holotype: Guinea, [Guinée française], Conakry, nov. 1930, A. Chevalier s.n. (PC 0789067 mixed with F. inaequalilimbatus; isotypes: PC0731291 and 0731292).
Diagnosis
The short, wide limbidial cells of the dorsal and apical lamina are distinctive of this species and differentiate it from the other species in this paper.
Etymology
The species has been named for its wide limbidium and limbidial cells (latus: wide).
Description
Brown in KOH. Stem 10–15 × 1.5–2 mm, branched or unbranched, pinnately foliated, hardly heterocaulous, with central strand; rhizoids brown, smooth; axillary nodules not differentiated; leaves close, up to 32 pairs, hardly crispate when dry, undulate when wet (undulation particularly manifest in KOH), lanceolate with acute apex, 1.4–1.6 × 0.3–0.4 (–0.5) mm, 3.5–4.5 times as long as wide, margin crenulate where elimbate, limbidia present on all laminae, less often restricted to the vaginant laminae, limbidia of dorsal and apical lamina irregular, often interrupted and composed of short wide cells (Fig. 5H), limbidia ending below the leaf apex and above the insertion of the dorsal lamina, confluent at the apex of the vaginant laminae, reaching the insertion of the vaginant laminae, on mid dorsal lamina 0–20 µm (0–3 cells) wide, 1–2-stratose; on the vaginant lamina 16–24 µm (3–cells) wide, 2–3 stratose; vaginant laminae ± 2/3 the leaf length, at base about as wide as stem, rounded at insertion, margins plane, subequal; dorsal lamina rounded at insertion, reaching the insertion, not decurrent; dorsal and apical lamina unistratose; costa ending 1 cell below the apex to per- or excurrent, in cross-section bryoides-type; mid dorsal laminal cells (4.0–) 5.5–7.0 × 3.0–5.0 (–6.5) µm, inconspicuously papillose; mid vaginant laminal cells (4.5–) 6.0–9.5 (–11.0) × 3.5–6.5 (–7.5) µm, pluripapillose, papillae in some leaves tiny and hard to observe, in others larger and clearly marginal (Fig. 5G) with 4–6 papillae per cell. Gemmae not observed. Perichaetia terminal, perichaetial leaves 1.5 mm long; perigonia, archegonia and sporophyte not observed.
Substrate and habit
Substrate, habitat and elevation not indicated. Growing in mats.
Notes
Three of the six acknowledged limbate and pluripapillose species, viz. F. weirii, F. dasyphus and F. latelimbatus are known to have limbate as well as semilimbate and transitional expressions. This suggests that the trait ‘having a limbidium on all laminae’ is less stable in this group than in subg. Fissidens where semilimbate expressions are much rarer.
Distribution
Guinea, known from the type only.
Mammillose species
Fissidens biformis Mitt. (Fig. 6)
J. Proc. Linn. Soc. Bot. 2: 141. 1859 – type: Sri Lanka [Ceylon], ad Matale, Gardner 626 -lectotype (designated by Bruggeman-Nannenga 2016): Ceylon, ad Matale, Gardner 55 (626) (NY 00966583; isolectotype: NY 00966584).
Fissidens angustifolius Sull., Proc. Am. Acad. Arts 5: 275. 1861 [1862] – lectotype (designated by Pursell 2007): CUBA, wet places on the ground in dense woods, Wright, distributed as Sullivant, Musci Cubensis 18 (FH; isolectotypes BM, NY; PC; the specimen in L contains F. zollingeri Mont.).
Fissidens brotheri Paris, Index Bryol. 462. 1896 (new name for F. leptophyllus Broth., Bot. Jahrb. Syst. 20: 182, 1895, hom. illeg.) – protologue: two syntypes: Togoland: Oiba und Ketschenké, auf Erdboden, Büttner s.n. – lectotype (designated by Bruggeman-Nannenga 1997): Togo, Oiba, ‘illegible’, 7-bri 1891, Büttner s.n. (H-BR).
Fissidens monostictus Broth. & P. de la Varde in Potier de la Varde, Bull. Soc. Bot France 72: 791, f. 19, 1a–c, 1925 – protologue: [Central African Republic], Yalinga. Ad terram. Le Testu 4808 – lectotype (designated by Pursell et al. 1992): Le Testu 4808A (MNHN-PC-PC0106471; isolectotypes: H-BR (Le Testu s.n.) and PC).
Diagnosis
This wide-spread variable species is characterized by small, frondose plants, completely limbate leaves, laminal cells distinctly to inconspicuously mammillose, basal juxta-costal vaginant laminal cells large (remark below), axillary nodules
present and peristome scariosus type. It resembles F. zollingeri Mont. in the mostly frondiform stems, limbate leaves, limbidia reaching the leaf apex and confluent with the costa, large vaginant laminal cells, axillary nodules and scariosus type peristomes. The two are distinct in the laminal cells that are smooth in F. zollingeri and mammillose in F. biformis. Furthermore, it can be confused with F. loennbergii P. de la Varde 1947 which also has frondose stems, well developed axillary nodules, limbate leaves and mammillose laminal cells. This species, however, has limbidia that end well below the leaf apex, shorter costae and its juxta-costal cells are not strongly inflated. Fissidens unipapillosus also has limbate leaves and mammillose cells, but otherwise is quite different for instance in having pinnately foliated plants.
Description
based on Arts REU 180/12 and Tisserant s.n., 25 X 1935)
Stem pale green, probably heterocaulous (vegetative stems not seen), 2.0 ×1.5 mm, unbranched or branched, frondosely foliated with 5–6 leaf pairs, central strand lacking; rhizoids pale brown, smooth; axillary nodules differentiated; leaves imbricate, rarely distant, green, crispate when dry, elliptico-falcate, with acute-acuminate apex, mid leaves 1.0–1.2 × 0.2–0.3 mm, 4–5 times as long as wide, limbate on all laminae of all well developed leaves; limbidium reaching the leaf apex and the insertions of dorsal and vaginant lamina, confluent at junction of vaginant laminae, in mid dorsal lamina 11 µm wide and (1–) 2-stratose, on vaginant laminae up to 24 µm wide, marginal to weakly intramarginal near the insertion, 1(–2)-stratose; vaginant laminae 3/5 leaf length, at the base less wide than the stem, slightly rounded at the insertion, subequal, unistratose; dorsal lamina tapering and straight towards insertion, reaching insertion, not decurrent; dorsal and apical lamina unistratose; costa excurrent, in cross -section bryoides-type; mid dorsal laminal cells (7.0–) 9.5–13.0 (–15.5) × 6.0–8.5 (–9.5) µm, distinctly mammillose; mid vaginant laminal cells (14.5–) 19.5–33.5 (–37.5) × (7.0–) 9.0–13.5, mammillose; basal juxta-costal cells of the vaginant lamina inflated, 39–62 × 12–24 µm. Gemmae not observed in African specimens, but reported by Pursell (2007). Perigonia and perichaetia terminal on different stems, perigonia stems short, 1.5 mm long; antheridia 180 µm long; archegonia 230–300 µm long; perichaetial leaves 1.7–1.9 mm long; calyptra weakly scabrous (Pursell 2007: smooth), 0.85 mm long. Sporophyte (only immature ones seen), seta 2–3 mm long, smooth; capsule too young for measurements; operculum (immature) rostrate, 0.6 mm long; spores not observed (10–13 µm and finely papillose according to Pursell 2007).
Notes
The term ‘large inflated juxta-costal cells’ was introduced by Pursell (1988) who was struck by the resemblance between the basal juxta-costal cells of F. zollingeri and F. biformis and the large, inflated cells of the species of subg. Polypodiopsis sect. Polypodiopsis and sect. Areofissidens Müll. Hal., 1848. He assumed that this resemblance indicates a close relationship between these taxa. This is supported by the peristome type and number of columns of exothecial cells (Bruggeman-Nannenga 2021). These large basal juxta-costal vaginant laminal cells are considered an important diagnostic character for F. zollingeri and F. biformis (Pursell 2007, Bruggeman-Nannenga 2017). Indeed, when present, such cells are diagnostic. Unfortunately, juxta-costal cells in these species are frequently only slightly enlarged, while Fissidens species from all subgenera frequently have more or less enlarged basal juxta-costal cells. Fissidens zollingeri and F. biformis with ‘ambiguous’ juxta costal cells can be confused with species of subg. Fissidens. When sporophytes are present, they are easily separated by the lower number of exothecial cells (± 32) and the scariosus peristome type (Bruggeman-Nannenga 2021) in F. biformis and F. zollingeri.
Substrate and habit
In Africa known from soil, less often on rocks (all collections from tree ferns cited by Bruggeman-Nannenga and Arts (2010) appeared to be F. loennbergii). In the neotropics known from soil, limestone rocks and decaying logs (Pursell 2007). Growing in tufts in forest, near a rivulet. Altitude: sea-level; (Pursell 2007: 0–1530 m).
Distribution
Pantropical. South and Central America, West Indies and some Pacific Islands. Uncommon in Africa. West Tropical Africa (Guinea, Nigeria (Egunyomi 1981), Togo); West-Central Tropical Africa (Central African Republic) and Western Indian Ocean: Madagascar, La Réunion) – New to Guinea. Also reported from Uganda (O'Shea et al. 2003, Bruggeman-Nannenga 2006a), but these specimens appeared to be F. loennbergii (Bruggeman-Nannenga 2007).
Selected specimens
Central African Republic [Oubangui], route Bocaranga à Kouroussa, 25 X 1935, Tisserant s.n. (PC-PV); – Guinea, Massif de Simandou. Lisowski 6560 pp (KRAM-B, 2 stems); – Madagaskar, Nosy BÉ [Nossi-bé], April 1879, Hildebrandt s.n. (L); – La Réunion: Petit Saint-Pierre. Source de Petit Saint-Pierre, 55°45′E, 21°04′S, Arts RÉU 180/12 (BR); – Togo, Ketschenkimay (orthographic error for Ketschenké?), 12 V 1891, Büttner s.n (H-BR); Oiba, Büttner s.n. (H-BR, lectotype F. brotheri Paris).
Illustrations
Bruggeman-Nannenga and Arts 2010: Fig. 19 (as F. angustifolius); Potier de la Varde 1925, Fig. 19, 1a–c (as F. monostictus); Pursell 1994: Fig. 52 (as F. angustifolius) and 2007: Fig. 123a–k (as F. angustifolius).
Figure 5.
Fissideens latelimbatus. (A) vegetative stem (colored by KOH), (B) perichaetial stem (colored by KOH), (C) leaves showing differences between limbidia of F. latelimbatus (C1) and F. dasyphus (C2), (D) leaf apex, (E) mid leaf, (F) basal part of leaf (colored by KOH), (G) papillae of mid vaginant laminal cells (colored by KOH), (H) short wide limbidial cells in mid dorsal lamina, (I) cross-section stem. All, except C2 from type F. latelimbatus, C2 from Dusén 906 (F. dasyphus).
Figure 6.
Fissidens biformis. (A) perichaetial stem, (B) perichaetial and small perigonial stem, (C) sporophytic stem, (D) leaves, (E) leaf, (F) leaf apex, (G) mid leaf, (H) insertion of leaf with axillary nodule (an), (I) cross-section the showing bryoides type costa and inflated cells of the vaginant lamina, (J) cross-section dorsal lamina. (A, D, F–J) from Arts RÉU 180/1, (B), (E) from type of F. leptophyllus Broth. and (C) from Kouroussa, Tisserant s.n.
Fissidens loennbergii P. de la Varde (Fig. 7)
Svensk Bot. Tidskr. 41: 3, f, 1, 1947. Holotype: Kenya, [Afr. or. Mt Kenia], 1911, E. Lönnberg (S).
Diagnosis
Fissidens loennbergii is characterized by frondose stems with well developed axillary nodules (Fig. 7E, K), mammillose laminal cells, limbidia on all lamina of at least upper and mid leaves, not reaching the leaf apex and ending above the insertion of the dorsal lamina. It resembles F. biformis. Both have frondiform plants, well developed axillary nodules, limbidia on all laminae and mammillose laminal cells. They can be distinguished by the longer limbidia and large inflated juxta-costal vaginant laminal cells of F. biformis. Limbidia of F. loennbergii end well below the leaf apex and its juxta-costal vaginant laminal cells are less strongly enlarged (compare Fig. 6H–7k). The Asian species F. benitotanii is similar, but lacks axillary nodules. The mammillose and completely limbate Polynesian F. raiatensis is easily distinguished from F. loennbergii by its wider, 1–3 stratose limbidia and leaves with small, scattered bistratose areas.
Description
Stems 2.0–4.5 × 1.5–3.0 mm, frondose, unbranched or branched, central strand absent to weak, not heterocaulous, hardly coloring in KOH; rhizoids brown, smooth; axillary nodules well differentiated; leaves green, close, 6–8 leaf pairs, not to slightly crispate when dry, linear, apex narrowly acute to acute-acuminate, upper leaves 1.0–2.5 × 0.1–0.4 mm, (4.1–) 4.8 –8.6 (–9.6) times as long as wide, margin serrulate near the apex, limbidium present on all laminae of at least the mid and upper leaves, in lower leaves present on all laminae to restricted to the vaginant lamina, ending below the leaf apex, reaching the insertion of the dorsal laminae to ending more or less far above insertion, in mid dorsal lamina 6.0–14.5 µm wide, uni- to bistratose, marginal; at the apex of the vaginant laminae confluent or almost confluent, becoming weak towards the insertion, on the vaginant lamina 24–41 µm wide, uni- to bistratose, marginal or intramarginal; vaginant laminae 1/2–3/5 the leaf length, at base less wide than the stem, almost straight towards the insertion, unistratose, subequal to slightly inequal; dorsal lamina tapering towards the insertion, reaching insertion, not decurrent; dorsal and apical lamina unistratose; costa ending 1–3 cells below apex to per- or excurrent, in cross-section bryoides-type; mid dorsal laminal cells (4.0–) 7.0–10.0 (–11.5) × (3.5–) 5.0–7.5 (–9.0) µm, mammillose; mid vaginant laminal cells (4.5–) 7.5–11.0 (–12.0) × 4.0–8.0 (–9.5) µm, mammillose; juxta-costal cells of vaginant lamina 13.0–20.0 (–25.0) × (9.7–) 11.0–16.5 (–18.0) µm. Gemmae not observed. Perigonia and perichaetia on the same as well as on separate stems; gametangia pure male, pure female and synoicous; perigonia on short, 0.25–0.35 mm long, budlike branches and terminal on up to 0.6 mm long, bud-shaped stems, rarely antheridia naked in the upper leaf axils; antheridia 130–160 µm long; perichaetia terminal on stems or, less often, branches; archegonia 260–300 µm long, perichaetial leaves ± 2.5 mm long; calyptra scabrous, 0.5 mm long. Sporophyte, seta 2.5–4.5 mm long, smooth; capsule cylindrical to gibbose, 0.6–0.8 × 0.2–0.3 mm, ± 32 columns of oblong exothecial cells; peristome scariosus-type, tooth base 32–48 µm; operculum rostrate, 0.5 mm long; spores subglobose, 8–11 × 7–10 µm diam., delicately papillose to smooth.
Substrate and habit
Typically lignicolous, mostly on tree ferns, also on rotting wood, less often rupicolous. Growing in loose mats or scattered between liverworts. Often growing together with F. jonesii Bizot. Growing in forests (montane, rain, fog, mossy etc.). Altitude: 1250–1960 m (550–1900 m according to Bruggeman-Nannenga (2007), but the 550 m is an orthographic error).
Distribution
Endemic to East Tropical Africa: Kenya, Tanzania and Uganda. Widespread in Tanzania.
Selected specimens
Kenya: [Afr. or.], Mt Kenia, 1911, Lönnberg s.n. (S); – Tanzania, Nguru Mts in Morogoro District. Along the path between Maskati and Nkombola villages, Kis & Pócs 9126/R (EGR, L); North Kilimanjaro, above Rongai Forest Station, Kimengelia Stream, Pócs 90134/E, a few stems intermixed with F. crispus Mont. (EGR, L); West Usambara Mts, bei Sakala, Engler 1021 (H-BR); on Gonja Hill E of Mgawahi village, Borhidi, Sebsebe, Hedrén and Iversen 8432/O, P (EGR); Shume-Magamba F. R., E of Shume village, near sawmill. Pócs 8447/G and 8447/AF (both EGR, L); on summit of Gonja peak E of Mgwashi village, Pócs 8505/Q (EGR); Balangai West F. R., forest on the E ridge and N slope of Kilimandege. Pócs 8507/J (EGR), K and Q (both EGR, L); Lushoto Distr., University of Dar es Salaam F. R., Mazumbai, Townsend 75/548 (E); – Uganda, Bushenyi. Kalinzu Forest Reserve near Nkombe sawmill. 0°23′S, 30°5′E, 4 II 1887, Hodgetts U 4552C (E, L), O'Shea U 2978A (E, L) and Porley 575A (E, L).
Illustrations
Bruggeman-Nannenga 2006a: Fig. 2 as F. angustifolius; Potier de la Varde 1947: f. 1.
Figure 7.
Fissidens loennbergii. (A–B) gonioautoicous stems, (A) in polarized light to show the limbidia, (C) sporophytic stem, (D) perigonial plant, (E) fertile stem with terminal perichaetium and perigonial and perichaetial branches, (F) leaf with perigonial bud, (G–H) leaf apices, (I) mid leaf, (J) basal part of leaf, (K) basal part of leaf with axillary nodule. (A–B, D–G) and (J) from Kis & Pócs 9126/R, (C) from Borhidi et al. 8432/O, P, (H–I) and K from Pócs 8507/K.
Figure 8.
Fissidens unipapillosus. (A) perichaetial (left) and vegetative stem, (B) leaves, (C) upper part of perichaetial stem with terminal archegonia, four naked axillary antheridia and one naked axillary archegonium, (D) leaf apex, (E) mid leaf, (F) leaf insertion, (G) mammillae in side view (vaginant lamina). All from isolectotype Wigginton U 3106A.
Fissidens unipapillosus Brugg.-Nann. (Fig. 8)
Cryptog., Bryol. 25 (4): 309, f. 1, 2004 – holotype: Uganda, Masindi District, Budongo Forest, Royal Mile, 1°42′N, 031°32′E, on soil deposited on log. 1060 m. 24 I 1997, M. J. Wigginton U 3106A (E; isotype L).
Fissidens glauculus var. gymnandrus P. de la Varde, Rev. Bryol., n.s 4 (2): 60, pl. 2, 1, 1931 – lectotype (holotype?) (designated here): Gabon, rochers de la Lipopa. Le Testu 6840 (PC-PV). syn. nov.
Diagnosis
This species is characterized by pinnately foliated stems, limbate leaves and mammillose cells. It resembles the pluripapillose F. dasyphus from which differs primarily by mammillose cells and the occurrence of naked axillary antheridia and archegonia. More material is needed to confirm the constancy of second trait. From the other two African limbate, mammillose species, F. biformis and F. loennbergii, it differs by the pinnately foliated stem, smaller cells (dorsal lamina cells 3.5–5.0 µm versus (4.0–)7.0–10.0 (–11.5) µm in F. loennbergii and (7.0–) 9.5–13.0(–15.5) µm in F. biformis and the wider leaf apices.
Description
Stem 4.0–7.0 × 2.2 mm, unbranched, pinnately foliated with up to 20 pairs of leaves, with central strand; rhizoids pale brown, smooth; axillary nodules weakly differentiated; leaves green, crispate when dry, undulate when wet, oblong with widely acute to rounded obtuse, often apiculate apex, 1.30–1.80 × 0.30–0.35 mm, 4.5–5.0 times as long as wide, margin denticulate near apex and base of dorsal lamina, limbate on all laminae of all or most leaves; limbidium marginal throughout, ending below the apex and well above insertion of dorsal lamina, confluent at junction of vaginant lamina, reaching the insertion of vaginant lamina, in mid dorsal lamina 9 µm wide, on the vaginant lamina up to 10.5 µm (three cells) wide, bistratose; vaginant laminae 3/4 leaf length, at base about as wide as stem, rounded at insertion, margins reflexed, unistratose, slightly unequal, strongly unequal in some perichaetial leaves; dorsal lamina rounded below, reaching the insertion, not decurrent; dorsal and apical lamina unistratose; costa per- to excurrent, in cross-section bryoides-type; laminal cells uniform throughout, roundish-quadrate, mid dorsal laminal cells 3.5–5.0 × 2.0–3.5 (–4.5) µm, highly and sharply mammillose (Fig. 8G), infrequently bipapillose; mid vaginant laminal cells 3.5–5.5 (–6.5) × 2.0–4.5 µm, highly and sharply mammillose, infrequently bipapillose. No gemmae observed. Perigonia and perichaetia on the same as well as on separate plants; antheridia axillary, on bud shaped perigonial branches or naked, solitary or in pairs (Fig. 8C); antheridia 120–200 µm long; perichaetia terminal, rarely axillary (Fig. 8C); perichaetial leaves 1.4–1.7 mm long, archegonia 200–300 µm long; calyptra scabrous, 0.55 mm long. Sporophyte, seta 3 mm long, smooth, capsule inclined, 0.70 × 0.35 mm, with ± 32 columns of quadrate, collenchymatous exothecial cells; peristome scariosus-type, basal inner peristome lamellae fimbriate, tooth base 45 µm wide; operculum rostrate, 0.5 mm long; spores 9.0–13.5 µm diam., smooth.
Substrate and habit
Mostly on soil, rarely on rocks, termite mounds or wood. Growing in mats. Growing in rainforest, damp places, mostly not indicated. Altitude: 1060–1220 m.
Distribution
Wide-spread, rare African endemic known from Gabon and Uganda. New to Gabon under this name.
Examined specimens
Gabon, rochers de la Lipopa, Le Testu 6840 (type of F. glauculus var. gymnandrus) (PC); – Uganda, Kampala District, Mpanga Forest Reserve, 3 VIII 1955, Jones 580 (paratype E, BM); Jones 603A (paratype BM), Mulange, 19 IV 1919, Dummer 4009 (paratype BM); West Nile District, Budonga forest, 15 VII1953, Chanchellor 13 (paratype BM); Damba Island II, Wood s.n. (paratype BM).
Illustrations
Bruggeman-Nannenga 2004: Fig. 1; Potier de la Varde 1931: pl. 2 and 1936: Fig. 7 as F. glauculus var. gymnandrus (same figure as in Potier de la Varde 1931).
Acknowledgements
I am greatly obliged to Sébastien Leblond who assembled relevant specimens of F. latelimbatus from the herbarium (PC), John Brinda for help with nomenclatorial questions, to the curators of BM, PC, EGR, Tamas Pócs, Ryzsard Ochyra and Gabriella Kis for the loan/gift of specimens and, last but not least, to Nicolien Sol (L) for manifold help.
© 2024 The Authors. This is an Open Access article
